Temporal variation may have diverse impacts on range limits.

Environmental fluctuations are pervasive in nature, but the influence of non-directional temporal variation on range limits has received scant attention. We synthesize insights from the literature and use simple models to make conceptual points about the potentially wide range of ecological and evolutionary effects of temporal variation on range limits. Because organisms respond nonlinearly to environmental conditions, temporal variation can directionally alter long-term growth rates, either to shrink or to expand ranges. We illustrate this diversity of outcomes with a model of competition along a mortality gradient. Temporal variation can permit transitions between alternative states, potentially facilitating range expansion. We show this for variation in dispersal, using simple source-sink population models (with strong Allee effects, or with gene flow hampering local adaptation). Temporal variation enhances extinction risk owing to demographic stochasticity, rare events, and loss of genetic variation, all tending to shrink ranges. However, specific adaptations to exploit variation (including dispersal) may permit larger ranges than in similar but constant environments. Grappling with temporal variation is essential both to understand eco-evolutionary dynamics at range limits and to guide conservation and management strategies. This article is part of the theme issue 'Species' ranges in the face of changing environments (Part II)'.

Disturbance-induced emigration: an overlooked mechanism that reduces metapopulation extinction risk.

Emigration propensity (i.e., the tendency to leave undisturbed patches) is a key life-history trait of organisms in metapopulations with local extinctions and colonizations. Metapopulation models of dispersal evolution typically assume that patch disturbance kills all individuals within the patch, thus causing local extinction. However, individuals may instead be able to leave a patch when it is disturbed, either by fleeing before being killed or simply because the disturbance destroys the patch without causing mortality. This scenario may pertain to a wide range of organisms from horizontally transmitted symbionts, to aquatic insects inhabiting temporary ponds, to vertebrates living in fragmented forests. We generalized a Levins-type metapopulation model of dispersal evolution by adding a new parameter of disturbance escape probability, which incorporates a second source of dispersal into the model: disturbance-induced emigration. We show that disturbance escape expands the domain of metapopulation viability and selects for lower rates of emigration propensity when disturbance rates are high. The fitness gains from disturbance-induced emigration are generally moderate, suggesting that disturbance escape might act more as a complementary dispersal strategy rather than a replacement to emigration propensity, at least for metapopulations that meet the assumptions of the Levins-type model. Yet disturbance-induced emigration may in some circumstances rescue a metapopulation from long-term extinction when the combination of high disturbance rates and low local population growth rates compromises its viability. Further, a metapopulation could persist exclusively by disturbance escape if local carrying capacities are large enough to counterbalance two sources of mortality: mortality driven by disturbance and mortality during dispersal. This study opens two promising research lines: (1) the investigation of disturbance escape in metapopulations of ephemeral habitats with unsaturated populations and non-equilibrium dynamics and (2) the incorporation of information costs to investigate the joint evolution of disturbance escape and emigration propensity.

Environmental fluctuations dampen the effects of clonal reproduction on evolutionary rescue.

Evolutionary rescue occurs when genetic change allows a population to persist in response to an environmental change that would otherwise have led to extinction. Most studies of evolutionary rescue assume that species have either fully clonal or fully sexual reproduction; however, many species have partially clonal reproductive strategies in which they reproduce both clonally and sexually. Furthermore, the few evolutionary rescue studies that have evaluated partially clonal reproduction did not consider fluctuations in the environment, which are nearly ubiquitous in nature. Here, we use individual-based simulations to investigate how environmental fluctuations (either uncorrelated or positively autocorrelated) influence the effect of clonality on evolutionary rescue. We show that, for moderate magnitudes of environmental fluctuations, as was found in the absence of fluctuations, increasing the degree of clonality increases the probability of population persistence in response to an abrupt environmental change, but decreases persistence in response to a continuous, directional environmental change. However, with large magnitudes of fluctuations, both the benefits of clonality following a step change and the detrimental effects of clonality following a continuous, directional change are generally reduced; in fact, in the latter scenario, increasing clonality can even become beneficial if environmental fluctuations are autocorrelated. We also show that increased generational overlap dampens the effects of environmental fluctuations. Overall, we demonstrate that understanding the evolutionary rescue of partially clonal organisms requires not only knowledge of the species life history and the type of environmental change, but also an understanding of the magnitude and autocorrelation of environmental fluctuations.

Threshold assessment, categorical perception, and the evolution of reliable signaling.

Animals often use assessment signals to communicate information about their quality to a variety of receivers, including potential mates, competitors, and predators. But what maintains reliable signaling and prevents signalers from signaling a better quality than they actually have? Previous work has shown that reliable signaling can be maintained if signalers pay fitness costs for signaling at different intensities and these costs are greater for lower quality individuals than higher quality ones. Models supporting this idea typically assume that continuous variation in signal intensity is perceived as such by receivers. In many organisms, however, receivers have threshold responses to signals, in which they respond to a signal if it is above a threshold value and do not respond if the signal is below the threshold value. Here, we use both analytical and individual-based models to investigate how such threshold responses affect the reliability of assessment signals. We show that reliable signaling systems can break down when receivers have an invariant threshold response, but reliable signaling can be rescued if there is variation among receivers in the location of their threshold boundary. Our models provide an important step toward understanding signal evolution when receivers have threshold responses to continuous signal variation.

Environmental fluctuations can promote evolutionary rescue in high-extinction-risk scenarios.

Substantial environmental change can force a population onto a path towards extinction, but under some conditions, adaptation by natural selection can rescue the population and allow it to persist. This process, known as evolutionary rescue, is believed to be less likely to occur with greater magnitudes of random environmental fluctuations because environmental variation decreases expected population size, increases variance in population size and increases evolutionary lag. However, previous studies of evolutionary rescue in fluctuating environments have only considered scenarios in which evolutionary rescue was likely to occur. We extend these studies to assess how baseline extinction risk (which we manipulated via changes in the initial population size, degree of environmental change or mutation rate) influences the effects of environmental variation on evolutionary rescue following an abrupt environmental change. Using a combination of analytical models and stochastic simulations, we show that autocorrelated environmental variation hinders evolutionary rescue in low-extinction-risk scenarios but facilitates rescue in high-risk scenarios. In these high-risk cases, the chance of a run of good years counteracts the otherwise negative effects of environmental variation on evolutionary demography. These findings can inform the development of effective conservation practices that consider evolutionary responses to abrupt environmental changes.

Digest: Propagule pressure might not matter in the establishment of invasive prokaryotes.

Lavigne et al. developed models to investigate the adaptive colonization of sink environments by asexual organisms. Their results have clear relevance to the spread of infectious disease, but they may also provide insights into prokaryotic invasions into natural communities. Their results show that propagule pressure might not be a good predictor of invasion success in prokaryotes, suggesting that more work is needed to understand how microbial invasions differ from those of plants and animals.

Pulsed Immigration Events Can Facilitate Adaptation to Harsh Sink Environments.

In nature, rates of dispersal vary greatly over time, yet most theoretical explorations of ecological and evolutionary dynamics to date have assumed constant movement rates. Here we examine how a particular pattern of temporal variation-periodic pulses of immigration-influences adaptation to a harsh environment, in which a species experiences conditions outside its niche requirements. Using both deterministic models and stochastic individual-based simulations, we show that for many ecological and genetic scenarios, temporally spacing out immigration events increases the probability that local adaptation is sufficient for persistence (i.e., niche evolution). When immigration events are too frequent, gene flow can hamper local adaptation in sexual species, but sufficiently infrequent pulses of immigration allow for repeated opportunities for adaptation with temporary escapes from gene flow during which local selection is unleashed. We develop versions of our models with and without density dependence for three different assumptions about the genetics underlying fitness (haploid, diploid, and quantitative genetic variation) so that our results may be applicable to a wide range of natural systems. Our study adds to a growing body of literature showing that temporal variation in migration rates can have significant effects on local adaptation and is among the first to show how such variation affects niche evolution.